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PubMed2000

Modulation of audiogenic seizures by histamine and adenosine receptors in the inferior colliculus.

Susceptibility to behaviorally similar audiogenic seizures (AGS) occurs genetically and is inducible during ethanol withdrawal (ETX). Comparisons between AGS mechanisms of genetically epilepsy-prone rats (GEPR-9s) and ethanol-withdrawn rats (ETX-Rs) are yielding information about general pathophysiological mechanisms of epileptogenesis. The inferior colliculus (IC) is the AGS initiation site. Excitatory amino acid (EAA) abnormalities in the IC are implicated in AGS, and histamine and adenosine receptor activation each reduce EAA release and inhibit several seizure types. Previous studies indicate that focal infusion of an adenosine receptor agonist into the IC blocked AGS in GEPR-9s, but the effects of adenosine receptor activation in the IC on AGS in ETX-Rs are unknown. The effects of histamine receptor activation on either form of AGS are also unexamined. The present study evaluated effects of histamine or a nonselective adenosine A(1) agonist, 2-chloroadenosine, on AGS by focal microinjection into the IC. Ethanol dependence and AGS susceptibility were induced in normal rats by intragastric ethanol. Histamine (40 or 60 nmol/side) significantly reduced AGS in GEPR-9s, but histamine in doses up to 120 nmol/side did not affect AGS in ETX-Rs. 2-Chloroadenosine (5 or 10 nmol/side) did not affect AGS in ETX-Rs, despite the effectiveness of lower doses of this agent in GEPR-9s reported previously. Thus, histamine and adenosine receptors in the IC modulate AGS of GEPR-9s, but do not modulate ETX-induced AGS. The reasons for this difference may involve the chronicity of AGS susceptibility in GEPR-9s, which may lead to more extensive neuromodulation as compensatory mechanisms to limit the seizures compared to the acute AGS of ETX-Rs.

2-Chloroadenosine↗
PubMed2000

Characterization of audiogenic-like seizures in naive rats evoked by activation of AMPA and NMDA receptors in the inferior colliculus.

The role of glutamate receptors in the inferior colliculus (IC) in audiogenic and audiogenic-like seizures was investigated in adult rats with transient neonatal hypothyroidism by 0.02% propylthiouracil (PTU) treatment through mother's milk (PTU rats) and in naive rats treated intracisternally with N-methyl-d-aspartate (NMDA), alpha-amino-3-hydroxy-5-methyl-4-isoxazole-proprionic acid (AMPA), or cyclothiazide, an inhibitor of rapid AMPA receptor desensitization. All rats showed audiogenic or audiogenic-like seizures characterized by running fit (RF) and generalized tonic-clonic seizures (GTCS). While systemically administered MK-801 inhibited GTCS, intracisternally administered NBQX inhibited RF and GTCS in both audiogenic and audiogenic-like seizures. Auditory stimulation shortened the latency to GTCS induced by AMPA, but not NMDA, at a subclinical dose and further elongated the shortened duration of RF, but not GTCS, induced by MK-801 pretreatment. Furthermore, Northern blot analysis was used to evaluate the expression of the immediate-early gene c-fos in the IC following induction of audiogenic or audiogenic-like seizures. The significant induction of c-fos mRNA by audiogenic seizures in PTU rats or by AMPA- or cyclothiazide-induced seizures in naive rats was prominent in the IC. MK-801 suppressed c-fos mRNA expression in the IC induced by audiogenic seizures in PTU rats or by AMPA-induced seizures in naive rats. NBQX suppressed the expression of c-fos mRNA in the IC induced by AMPA-induced seizures but did not suppress c-fos mRNA in PTU rats or rats with cyclothiazide-induced seizures. Auditory stimuli failed to affect c-fos mRNA induction by AMPA. The present study suggests that audiogenic-like seizures can be reproduced by glutamate receptor agonists in which AMPA receptors are primarily linked to the initiation of audiogenic seizures (RF) while NMDA receptors presumably located within the IC are involved in the propagation of GTCS in audiogenic seizures.

Acoustic Stimulation↗
PubMed2000

Tectal graft modulation of audiogenic seizures in Long-Evans rat.

Audiogenic seizure (AGS) activity can be induced in the seizure-resistant Long-Evans rat by postnatal priming. This study examined the effects of unilateral lesions of the inferior colliculus (IC) and implantation of tectal grafts on AGS components. Animals were primed with a 10-kHz tone burst at 120 dB on postnatal day 14 and tested for AGS susceptibility on day 28, and then two groups were unilaterally lesioned including animals receiving embryonic day 16-17 grafts of caudal tectum. Subsequently, animals were repeatedly tested for wild running and clonic-tonic convulsion components of AGS. The results demonstrate that unilaterally grafted animals with partial IC lesions showed significant reduction in the incidence of clonus expression with greater terminal uniphasic wild running behavior. These effects were stronger than in animals with comparable unilateral lesions alone. Many neurons in graft cases were in direct contact with host tissues to provide a substrate for tissue interactions previously demonstrated to promote neuron survival and remediate IC functions.

Acoustic Stimulation↗
PubMed2001

Audiogenic seizure susceptibility is induced by termination of continuous infusion of gamma-aminobutyric acid or an N-methyl-D-aspartic acid Antagonist into the inferior colliculus.

The inferior colliculus (IC) is strongly implicated in seizure initiation in a genetic form of audiogenic seizures (AGS) and in AGS observed during ethanol withdrawal (ETX). Ethanol is known to block the actions of excitatory amino acids (EAA) and enhance the actions of gamma-aminobutyric acid (GABA) in several brain areas, including the IC. The present study investigated the effects on susceptibility to AGS following withdrawal from continuous blockade of N-methyl-D-aspartic acid (NMDA) receptors or continuous activation of GABA receptors in the IC. This involved infusion of GABA (1 M) or a competitive NMDA antagonist, DL-2-amino-7-phosphonoheptanoic acid (AP7, 1 mM), at 0.25 microl/h for 7 days using an Alzet osmotic minipump. Following abrupt termination of the infusion, AGS susceptibility began at 30 min. The incidence of AGS was 38.9 and 56.3% following GABA and AP7 withdrawal, respectively. The AGS behaviors observed during withdrawal, which included wild running and bouncing clonus, were very similar to those evoked by acoustic stimuli during ETX. AGS susceptibility lasted for several hours and in 13% of animals persisted for up to 6 months. The current results support diminished GABAergic and elevated glutamatergic function in the IC as the critical mechanisms and sites for AGS initiation. The present study, coupled with previous evidence that chronic ethanol exposure reduced GABA-mediated inhibition and enhanced EAA-mediated excitation, suggests that these amino acid receptor-mediated alterations in the IC are key elements in initiating AGS during ethanol withdrawal.

2-Amino-5-phosphonovalerate↗
PubMed2002

Transplantation of M213-2O cells with enhanced GAD67 expression into the inferior colliculus alters audiogenic seizures.

The purpose of the present study was to examine the effects of GABA-producing cell transplants on audiogenic seizures (AGS). The M213-2O cell line was derived from fetal rat striatum and has GABAergic properties. This cell line was further modified to express human GAD(67) and produce elevated levels of GABA. The present study compares the effects of parent M213-2O cell transplants with those of GAD(67)-modified M213-2O cells in AGS-prone Long-Evans rats. Two weeks following implantation of engineered cells, latency to AGS-typical wild running was increased compared to nonimplanted subjects. Survival of the transplanted cells was confirmed by immunochemical labeling of GAD(67) and Epstein-Barr virus nuclear antigen. These findings support the use of GABA-producing cell lines to modify seizure activity.

Acoustic Stimulation↗
PubMed1997

Metabolic effects of blocking tone conditioning on the rat auditory system.

The Kamin blocking phenomenon occurs when behavioral expression of conditioning to a novel stimulus fails in the presence of a previously conditioned stimulus (CS). Neural metabolic effects of a tone conditioned as an excitor were compared to the effects of the same physical tone when excitatory conditioning was blocked by previous conditioning with a light. We examined the metabolic activity of the auditory system to test the hypothesis that auditory processing of a tone CS changes during blocking. Quantitative histochemistry of cytochrome oxidase (C.O.), the final mitochondrial enzyme for oxidative metabolism, was used to evaluate cumulative changes in the metabolic capacity of the auditory system resulting from blocking. Rats (Long-Evans) in the Blocking group received pairings of a light CS with a mild footshock unconditioned stimulus (US) during Phase 1 training. Rats in the Control group received random presentations of the same stimuli during Phase 1. Both groups then received the same Phase 2 training consisting of simultaneous tone and light presentations paired with footshock. The Control group exhibited significant suppression of drinking to tone alone presentations after training, whereas the Blocking group did not. Metabolic mapping results demonstrated that blocking effects were localized to auditory regions receiving direct US somatosensory projections. Significantly greater C.O. activity in the inferior colliculus and the dorsal cochlear nucleus was found for the Blocking group relative to the Control group. Input cell layers of secondary auditory cortex also demonstrated a group difference, in that layers II/III and IV had lower levels of C.O. activity in the Blocking group. These specific changes in C.O. activity linked to behavioral training demonstrated that the blocking phenomenon produced distinct neural metabolic changes in CS processing in the auditory system localized to regions with CS-US interactions.

Animals↗
PubMed1995

Expression of the c-fos transcription factor in the rat auditory pathway following postnatal auditory deprivation.

As an animal model for inborn hearing loss rat pups were reared in a sound-proof chamber from birth until age 21 days. In addition, pinnae were bilaterally sutured closed to reduce any influence of ambient sound. At the end of the sound deprivation, outer ear channels were reopened. Since previous studies failed to show any difference in the number or morphology of neurons in the auditory pathway in bilaterally sound-deprived animals, expression of c-fos protein was used as a functional marker to map trans-synaptic information transfer in the auditory pathway with cellular resolution. At day 21 sound-deprived rats and untreated controls were stimulated with pure tones of 8kHz for 5min at different sound pressure levels. Acoustic stimulation induced c-fos in both parts of the cochlear nucleus, superior olivary complex and inferior colliculus. Compared to untreated rats, deprivation reduced the number of c-fos labeled neurons in the dorsal and ventral part of the cochlear nucleus and inferior colliculus by 58% and 30%, respectively, following low sound pressure levels (90dB). In contrast, high sound pressure levels (120dB) increased the number of c-fos labeled neurons in these areas and evoked only minor differences in the number of labeled neurons in both untreated and sound deprived rats.

Acoustic Stimulation↗
PubMed1996

Distribution of calcium-binding protein immunoreactivities in the guinea pig auditory brainstem.

This study was intended to provide an overview of the distribution of calcium-binding proteins in the rodent auditory brainstem. We based our observations on immunohistochemical material obtained in the guinea pig, a species widely used in auditory research in which a mapping of calcium-binding proteins in the auditory brainstem is still missing. Differences in the amounts of these proteins throughout the auditory brainstem were further analyzed semiquantitatively. Parvalbumin was present in most neurons and their axon terminals throughout the ascending auditory brainstem. Nuclei that surround the main relay nuclei of the ascending auditory pathway lacked labeling. Calretinin staining was prominent in spherical and globular cells of the cochlear nucleus, in their axon terminals in the superior olivary complex, and in principal cells of the medial superior olive. Measures of optical densities showed that auditory neurons involved in sound localization had the highest calretinin labeling levels. Calbindin D-28k was present in cartwheel cells of the dorsal cochlear nucleus, in almost all neurons of the medial nucleus of the trapezoid body, and in globular cells in the ventral nucleus of the lateral lemniscus. The labeling patterns for calretinin and calbindin D-28k were non-overlapping throughout the auditory brainstem. This was also evident in the ventral nucleus of the lateral lemniscus where calbindin D-28k-immunoreactive terminals were found in the medial portion, while the calretinin-immunoreactive terminals were observed in the lateral portion. This study presents the first direct and comprehensive comparison of these three calcium-binding proteins in the auditory brainstem of a rodent. Each antibody yields a unique staining pattern that provides a basis for further defining neuronal populations. In addition, since their axons are also selectively stained, auditory nuclei can further be compartmentalized based on different terminal fields. These immunoreactivities have provided clues to the complex structure of the auditory brainstem.

Animals↗
PubMed1996

Neurons with different temporal firing patterns in the inferior colliculus of the little brown bat differentially process sinusoidal amplitude-modulated signals.

We examined how well single neurons in the inferior colliculus (IC) of an FM bat (Myotis lucifugus) processed simple tone bursts of different duration and sinusoidal amplitude-modulated (SAM) signals that approximated passively heard natural sounds. Units' responses to SAM tones, measured in terms of average spike count and firing synchrony to the modulation envelope, were plotted as a function of the modulation frequency to construct their modulation transfer functions. These functions were classified according to their shape (e.g., band-, low-, high-, and all-pass). IC neurons having different temporal firing patterns to simple tone bursts (tonic, chopper, onset-late, and onset-immediate) exhibited different selectivities for SAM signals. All tonic and 83% of chopper neurons responded robustly to SAM signals and displayed a variety of spike count-based response functions. These neurons showed a decreased level of time-locking as the modulation frequency was increased, and thereby gave low-pass synchronization-based response functions. In contrast, 64% of onset-immediate, 37% of onset-late and 17% of chopper units failed to respond to SAM signals at any modulation frequency tested (5-800 Hz). Those onset neurons that did respond to SAM showed poor time-locking (i.e., non-significant levels of synchronization). We obtained evidence that the poor SAM response of some onset and chopper neurons was due to a preference for short-duration signals. These data suggest that tonic and most chopper neurons are better-suited for the processing of long-duration SAM signals related to passive hearing, whereas onset neurons are better-suited for the processing of short, pulsatile signals such as those used in echolocation.

Animals↗
PubMed1990

Ontogenesis of auditory fovea representation in the inferior colliculus of the Sri Lankan rufous horseshoe bat, Rhinolophus rouxi.

This report describes the ontogenesis of tonotopy in the inferior colliculus (IC) of the rufous horseshoe bat (Rhinolophus rouxi). Horseshoe bats are deaf at birth, but consistent tonotopy with a low-to-high frequency gradient from dorsolateral to ventromedial develops from the 2nd up to the 5th week. The representation of the auditory fovea is established in ventro-medio-caudal parts of the IC during the 3rd postnatal week (Fig. 3). Then, a narrow frequency band 5 kHz in width, comprising 16% of the bat's auditory range, captures 50-60 vol% of the IC (Fig. 3c). However, foveal tuning is 10-12 kHz (1/3 octave) lower than in adults; foveal tuning in females (65-68 kHz) is 2-3 kHz higher than in males (62-65 Khz). Thereafter, foveal tuning increases by 1-1.5 kHz per day up to the 5th postnatal week, when the adult hearing range is established (Figs. 4, 5). The increase of sensitivity and of tuning sharpness of single units also follows a low-to-high frequency gradient (Fig. 6). Throughout this development the foveal tuning matches the second harmonic of the echolocation pulses vocalised by these young bats. The results confirm the hypothesis of developmental shifts in the frequency-place code for the foveal high frequency representation in the IC.

Acoustic Stimulation↗
PubMed1990

Audiovocal interactions during development? Vocalisation in deafened young horseshoe bats vs. audition in vocalisation-impaired bats.

Horseshoe bats (Rhinolophus rouxi) were deafened in their 3rd-5th postnatal week. Subsequently their vocalisations were monitored to evaluate the impact of audition on the development of echolocation pulses. Hearing impairment affected the echolocation pulses as follows: the frequency of the constant frequency (CF) component was altered by between +4 kHz and -14 kHz, and the dominance of the second harmonic of the pulses was neutralised by a relative increase in intensity of the first and third harmonics. A second experiment focused on possible influences of acoustical self-stimulation with echolocation pulses on the establishment of auditory fovea representation in the inferior colliculus (IC). Frequency control of echolocation pulses was disrupted by larynx denervation. Thereafter, the bats produced multiharmonic echolocation signals (4-11 harmonics) varying in frequency. IC tonotopy, however, as monitored by stereotaxic electrophysiology, showed the same developmental dynamics as seen in control specimens (Fig. 10). Both experiments indicate that throughout postnatal development echolocation pulses are under auditory feedback control, whereas maturation of the auditory fovea and shifts in its frequency tuning represent an innate process. The significance of this postnatal development might be the adjustment of the vocal motor system of each bat to the frequency of its 'personal' auditory fovea.

Animals↗
PubMed1996

Effects of sound direction on the processing of amplitude-modulated signals in the frog inferior colliculus.

Single-unit recordings were made from 143 neurons in the frog (Rana p. pipiens) inferior colliculus (IC) to investigate how free-field sound direction influenced neural responses to sinusoidal-amplitude-modulated (SAM) tone and/or noise. Modulation transfer functions (MTFs) were derived from 3 to 5 sound directions within 180 degrees of frontal field. Five classes of MTF were observed: low-pass, high-pass, band-pass, multi-pass, and all-pass. For 64% of IC neurons, the MTF class remained unchanged when sound direction was shifted from contralateral 90 degrees to ipsilateral 90 degrees. However, the MTFs of more than half of these neurons exhibited narrower bandwidths when the loudspeaker was shifted to ipsilateral azimuths. There was a decrease in the cut-off frequency for neurons possessing low-pass MTFs, an increase in cut-off frequency for neurons showing high-pass MTFs, or a reduction in the pass-band for neurons displaying bandpass MTFs. These results suggest that sound direction can influence amplitude modulation (AM) frequency tuning of single IC neurons. Since changes in periodicity of SAM tones alter both the temporal parameters of sounds as well as the sound spectrum, we examined whether directional effects on spectral selectivity play a role in shaping the observed direction-dependent AM selectivity. The directional influence on AM selectivity to both SAM tone and SAM noise was measured in 62 neurons in an attempt to gain some insight into the mechanisms that underlie directionally-induced changes in AM selectivity. Direction-dependent changes in the shapes of the tone and noise derived MTFs were different for the majority of IC neurons (55/62) tested. These data indicate that a spectrally-based and a temporally-based mechanism may be responsible for the observed results.

Acoustic Stimulation↗
PubMed1996

Responses to simulated echoes by neurons in the barn owl's auditory space map.

The natural acoustical environment contains many reflective surfaces that give rise to echoes, complicating the task of sound localization and identification. The barn owl (Tyto alba), as a nocturnal predator, relies heavily on its auditory system for tracking and capturing prey in this highly echoic environment. The external nucleus of the owl's inferior colliculus (ICx) contains a retina-like map of space composed of "space-specific" auditory neurons that have spatially limited receptive fields. We recorded extracellularly from individual space-specific neurons in an attempt to understand the pattern of activity across the ICx in response to a brief direct sound and a simulated echo. Space-specific neurons responded strongly to the direct sound, but their response to a simulated echo was suppressed, typically, if the echo arrived within 5 ms or less of the direct sound. Thus we expect there to be little or no representation within the ICx of echoes arriving within such short delays. Behavioral tests using the owl's natural tendency to turn their head toward a sound source suggested that owls, like their space-specific neurons, similarly localize only the first of two brief sounds. Naive, untrained owls were presented with a pair of sounds in rapid succession from two horizontally-separated speakers. With interstimulus delays of less than 10 ms, the owl consistently turned its head toward the leading speaker. Longer delays elicited head turns to either speaker with approximately equal frequency and in some cases to both speakers sequentially.

Acoustic Stimulation↗
PubMed1992

On the ability of neurons in the barn owl's inferior colliculus to sense brief appearances of interaural time difference.

This paper investigates the ability of neurons in the barn owl's (Tyto alba) inferior colliculus to sense brief appearances of interaural time difference (ITD), the main cue for azimuthal sound localization in this species. In the experiments, ITD-tuning was measured during presentation of a mask-probe-mask sequence. The probe consisted of a noise having a constant ITD, while the mask consisted of binaurally uncorrelated noise. Collicular neurons discriminated between the probe and masking noise by showing rapid changes from untuned to tuned and back to untuned responses. The curve describing the relation between probe duration and the degree of ITD-tuning resembled a leaky-integration process with a time constant of about 2 ms. Many neurons were ITD-tuned when probe duration was below 1 ms. These extremely short effective probe durations are interpreted as evidence for neuronal convergence within the pathway computing ITD. The minimal probe duration necessary for ITD-tuning was independent of the bandwidth of the neurons' frequency tuning and also of the best frequency of a neuron. Many narrowly tuned neurons having different best frequencies converge to form a broad-band neuron. To yield the short effective probe durations the convergence must occur in strong temporal synchronism.

Acoustic Stimulation↗
PubMed1992

Spatial processing within the mustache bat echolocation system: possible mechanisms for optimization.

1. The directionality of an echolocation system is determined by the acoustic properties of both the emitter and receiver, i.e., by the radiation pattern of the emitted pulse and the directionally of the external ears. We measured the directionality of the echolocation system of the greater mustache bat (Pteronotus parnellii) at the 30 kHz, 60 kHz and 90 kHz harmonics of its echolocation pulse by summing, at points throughout the frontal sound field, the echo attenuation due to the spread of pulse energy and the attenuation due to the spread of pulse energy and the attenuation due to the directionality of its external ears. The pulse radiation pattern at the 3 harmonics was measured by comparing the output of a microphone moved throughout the frontal sound field against a second reference microphone at the center of the field. External ear directionality at the 3. harmonics was measured by presenting free-field sounds throughout the frontal sound field, and recording the intensity thresholds of cochlear microphonic potentials, and the intensity thresholds of monaural neurons in the inferior colliculus tuned to one of the 3 harmonics. 2. When compared with ear directionality, the echolocation system was found to be more directional for the center of the sound field in several respects. At all harmonics, attenuation of sounds originating in the peripheral part of the field was increased by 10 to 13 dB. Areas of maximum sound intensity contracted toward the center of the field. Also, the isointensity contours of the echolocation system were more radially symmetrical about the center of the field. 3. At 60 kHz, sound intensity along the azimuth within the echolocation system was nearly constant 26 degrees to either side of the center of the field. This suggests that the radiation pattern of the echolocation pulse and the directionality of the external ears complement one another to produce an acoustic environment at the center of the sound field in which stimulus intensity is stabilized to allow more effective analysis of various aspects of the echolocation target. In particular, we suggest that this intensity stabilization may allow the bat to more effectively resolve the interaural intensity differences it uses to localize prey. 4. Predictions of the azimuthal spatial tuning of binaurally sensitive neurons in the inferior colliculus within the echolocation system were compared with their spatial tuning when only ear directionality is considered.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1994

Frequency tuning and response latencies at three levels in the brainstem of the echolocating bat, Eptesicus fuscus.

To determine the level at which certain response characteristics originate, we compared monaural auditory responses of neurons in ventral cochlear nucleus, nuclei of lateral lemniscus and inferior colliculus. Characteristics examined were sharpness of frequency tuning, latency variability for individual neurons and range of latencies across neurons. Exceptionally broad tuning curves were found in the nuclei of the lateral lemniscus, while exceptionally narrow tuning curves were found in the inferior colliculus. Neither specialized tuning characteristic was found in the ventral cochlear nuclei. All neurons in the columnar division of the ventral nucleus of the lateral lemniscus maintained low variability of latency over a broad range of stimulus conditions. Some neurons in the cochlear nucleus (12%) and some in the inferior colliculus (15%) had low variability in latency but only at best frequency. Range of latencies across neurons was small in the ventral cochlear nucleus (1.3-5.7 ms), intermediate in the nuclei of the lateral lemniscus (1.7-19.8 ms) and greatest in the inferior colliculus (2.9-42.0 ms). We conclude that, in the nuclei of the lateral lemniscus and in the inferior colliculus, unique tuning and timing properties are built up from ascending inputs.

Animals↗